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Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2015 | 7(6): 7253–7268 Review On the molluscan fauna of Lakshadweep included in various schedules of Wildlife (Protection) Act of India

A. Bijukumar 1, R. Ravinesh 2, A.R. Arathi 3 & K.K. Idreesbabu 4 ISSN 0974-7907 (Online) ISSN 0974-7893 (Print)

1,2,3 Department of Aquatic Biology and Fisheries, University of Kerala, Thiruvananthapuram, Kerala 695581, India 4 Department of Science and Technology, Kavaratti, U.T. of Lakshadweep 682555, India OPEN ACCESS 1 [email protected] (corresponding author),2 [email protected], 3 [email protected], 4 [email protected]

Abstract: Out of the 24 of marine molluscs included in Schedule I and IV of the Wildlife (Protection) Act (WPA) of India, 19 species were recorded from the coastal waters of Lakshadweep. A recent survey conducted by the authors recorded the presence of 14 scheduled molluscs in Lakshadweep. Scheduled species such as Placuna placenta (recorded from Kavaratti) and Tudicla spirillus (recorded from Kalpeni) are new records from Lakshadweep. The paper provides details for taxonomic identification of scheduled molluscs and discusses strategies for conservation of scheduled molluscs of Lakshadweep.

Keywords: Bivalvia, Cephalopoda, conservation, , , Nautilus, Wildlife (Protection) Act.

Malayalam Abstract: ഇn#യിൽ വന#ജീവി (സംരkണ) നിയമtിൽ പ"ിക I, IV എnിവയിൽ ഉൾെp! െമാtം 24 ൈജവജാതി കടൽ െമാള%കുകളിൽ 19 എ"ം ലkദ$ീപിെn തീരkടലിൽ നിn് കെ'tി. Placuna placenta (കവറtിയിൽ നിn് കെ'tിയ*), Tudicla spirillus (കൽേpനിയിൽ നിn് കെ'tിയ*) എnിവെയ ലkദ$ീപിൽ നിn് ആദ#മായാ' കെ*tുn.. പ0ികയിൽ െപ# ലkദ$ീപിെല െമാള"കുകെള തിരിcറിയാൻ സഹായിkുn വിശദവിവരŋള/ം അവയുെട സംരkണtിനുll ത:nŋള/ം ഈ േലഖനtിൽ വി#തരിc' !പതിപാദിkുnു.

Tridacna maxima

NOT DATA NEAR CRITICALLY EXTINCT LEAST VULNERABLE ENDANGERED EXTINCT EVALUATED DEFICIENT CONCERN THREATENED ENDANGERED IN THE WILD NE DD LC NT VU EN CR EW EX

DOI: http://dx.doi.org/10.11609/JoTT.o4140.7253-68 | ZooBank: urn:lsid:zoobank.org:pub:46141C50-3DFF-408E-A6B5-0963AF23AA7F

Editor: Anonymity requested. Date of publication: 26 May 2015 (online & print)

Manuscript details: Ms # o4140 | Received 02 September 2014 | Final received 30 April 2015 | Finally accepted 02 May 2015

Citation: Bijukumar, A., R. Ravinesh, A.R. Arathi & K.K. Idreesbabu (2015). On the molluscan fauna of Lakshadweep included in various schedules of Wildlife (Protection) Act of India. Journal of Threatened Taxa 7(6): 7253–7268; http://dx.doi.org/10.11609/JoTT.o4140.7253-68

Copyright: © Bijukumar et al. 2015. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use of this article in any medium, reproduc- tion and distribution by providing adequate credit to the authors and the source of publication.

Funding: Department of Biotechnology, Government of India (BT/PR13602/AAQ/03/508/2010).

Competing interests:The authors declare no competing interests.

For Author Details and Author Contributionsee end of this article.

Acknowledgements: The authors thank the Department of Biotechnology, Government of India for the financial support of the project (BT/PR13602/ AAQ/03/508/2010). The authors also thank the Department of Science and Technology (Permission F No. 42/4/2011-S&T dated 28.1.2014; F No. 42/4/2011- S&T (Vol.1)/210 dated 09.03.2015) and the Department of Environment and Forests, Union Territory of Lakshadweep (Permission F. No. 16/8/2008-E&F dated 7.5.2012 and 23.3.2013; F.No. 19/5/2012- E&F dated 19.3.2015) for the official permission to study marine molluscs of the islands. Arathi thanks the fellowship offered by the Kerala State Council for Science, Technology and Environment for the PhD programme. IdentificationTridacna of spp. was confirmed by Dr. Jan Johan ter Poorten, Zoological Museum, Amsterdam. Good quality photographs of indomaris were given by Mr. Ulrich Wieneke, Murnau, Germany. Thanks are due to M/s Sandy Beach (Kavaratti) for their help in field work.

7253 Molluscan fauna of Lakshadweep Bijukumar et al.

INTRODUCTION 2012; Susan et al. 2012; Venkataraman et al. 2012; Aditi & Apte 2013; Ravinesh et al. 2013; Vishal & Apte 2013; Considered as biological hotspots and the legacy of Apte & Vishal 2014). a distinctive evolutionary history, island biodiversity is Considering the increasing threats to marine unique owing to taxonomic distinctness, endemism and biodiversity, the Government of India has included specific requirements (da Fonseca et al. 2006). many marine species under the four schedules of Conversely, the risk of of species is much Wildlife (Protection) Act (WPA), 1972 (http://envfor.nic. more pronounced in islands as the populations are in/division/wildlife). Under Indian WPA the species to rather restricted, small and specific to certain . be protected are listed under Schedules I to VI. While Anthropogenic interventions have catastrophic effects Schedule I and II (Part II) species are covered by the most on island biodiversity and therefore demand urgent stringent regulations in the Act to get the highest level attention of conservationists and policy makers. The of legal protection, species in Schedules II (Part I), III and Secretariat of the Convention on Biological Diversity IV are given progressively narrow levels of protection; (CBD) considered the theme of the International Day Schedule V represents species labelled ‘vermin’ which for Biological Diversity (IDB) on 22 May 2014 as “Island are not afforded protection of any kind and Schedule Biodiversity”. In the case of Lakshadweep, the entire VI includes six (Prashanth & Veenakumari 1996; economy of the islands and the great cultural ethos of Kunte 2008). A total of 24 species of marine molluscs of the islanders remain closely interlinked with the rich India have been included under Schedule I and IV of WPA biodiversity. The threats to biodiversity are primarily (Ramakrishna & Dey 2003). This paper documents 19 due to unsustainable developmental activities, increase species of marine molluscs recorded from Lakshadweep, in human population, overexploitation and climate represented under various schedules of WPA, compiled change (KSCSTE 2013). from various databases/publications and also based on Mollusca comprise highly diverse phylum of the primary data collected. invertebrates, with accepted described living species of molluscs varying from 50,000 to a maximum of 1,20,000 species, and form the largest marine phylum, comprising MATERIALS AND METHODS about 23% of all the named marine organisms (Chapman 2009). They form an economically valuable and India’s smallest Union Territory Lakshadweep is are well represented in exports of marine products and an archipelago consisting of 36 islands with an area in domestic trade in the form of various products such of 32km2. It is a uni-district Union Territory and it as crafts, curios and . In tropical seas they occupy comprises 12 atolls, three reefs, five submerged banks every trophic level, ranging from primary producers to and 10 inhabited islands, which include Kavaratti, Agatti, top level carnivores (Mohamed 2012). The diversity of Amini, Androth, Kiltan, Kalpeni, Kadamat, Chetlat, Bitra marine molluscs in India is estimated as 3,370 species by and Minicoy (Fig. 1). It is located between 8–12 0N and Venkataraman & Wafar (2005) and as 2,300 by Tripathy 71–74 0E, 220–440 km away from the coastal city of & Mukhopadhyay (2015). The island ecosystems in Kochi in Kerala. The islands are ring-shaped atolls lying India are highly biodiverse with regard to molluscan along a north-south axis (except Androth) with a lagoon fauna. While the diversity of molluscan fauna of the to the west and open sea to the east. Lakshadweep is Andaman and Nicobar Islands includes 1,282 species the only coral atoll of the country. With a vast lagoon (Ramakrishna & Dey 2010), in Lakshadweep there are of 4,200km2, it has territorial waters of 20,000km2, an about 424 species (Rao & Rao 1991). A recent database Exclusive Economic Zone (EEZ) of 4,00,000km2 and a prepared by KSCSTE (2013) shows that the molluscan coast line of about 132km. Lakshadweep, Maldives and diversity in Lakshadweep includes 469 species. Surveys Chagos are three island groups in the Indian that on molluscan fauna of Lakshadweep by various authors together form a vast submarine mountain range, the in the last few decades are an indication of not having Chagos-Laccadive Plateau. This volcanic range lies just fully explored the molluscan diversity of the island east of the Mid-Indian Ridge and west of the Mid-Indian and therefore the number of molluscan species is Basin, and the chain of islands comprises the most probably underestimated (Namboodiri & Sivadas 1979; extensive and atoll community in the Indian Appukuttan et al. 1989; Apte 1998, 2009, 2012a,b, 2014; Ocean as well as the largest atoll system in the World Ramakrishna & Dey 2010; Rao 2003; Apte & Sutirtha (Olson & Dinerstein 1998). 2010; Apte et al. 2010, 2012, 2014; Prabhakaran et al. As part of the present study, surveys were carried out

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Figure 1. Lakshadweep islands

Figure 2. Generalised diagram of a bivalve mollusc in Agatti, Androth, Bangaram (uninhabited island), Bitra, photo documented using methods such as SCUBA diving, Kadamat, Kalpeni, Kavaratti and Minicoy islands for the snorkelling and intertidal handpicking. The photographs documentation of molluscs at multiple intervals from of scheduled molluscs and their life history stages were January 2012 to September 2014. The organisms were prepared based on specimens photographed in the

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1. chiragra (Linnaeus, 1758) (Images 1a-k) Appukuttan et al. 1989, p.79; Rao & Rao 1991, p. 277; Ramakrishna & Dey 2003, p 28; KSCSTE 2013, p. 146; Apte 1998, p.31; Apte 2014, p. 59. Description: Very thick, robust, ovate and heavy shell, with a distinct anterior notch. very obtuse and knotty. Six long and curved marginal digitations, expanded from the flaring, thick outer and canals (Image 1 i, j). Anterior canal and posterior most digitation of outer lip with their bases recurved at right angle to the left side. Deep between 4th and 5th labial digitations. Whorls 9–11; the four uppermost whorls are depressed, so as to give the spire a truncated appearance and each of the other whorls have a row of strong rounded knobs. Inner lip serrated. White exterior with brown patches and flecks; rose pink or orange interior. Columellar brown in specimens from Lakshadweep and chocolate brown, and are distinct from Pacific specimens (unpublished data). and aperture lirate (sculptured with fine lines or grooves). chitinous, dark brown, fusiform, with fine striations on one side. In the initial developmental phases, H. chiragra Figure 3. Generalised diagram of a gastropod mollusc shells have thin outer lips, without any fingers (Images 1a-d). As they grow, the outer lips become thick, expand and develop digits (Images 1 e-h). field or otherwise mentioned. Generalised diagrams Shell length 280–320 mm; Maximum length 320mm. of gastropod and bivalve molluscs are provided in Figs Remarks: In Lakshadweep it is found around all 2 and 3. inhabited islands (Appukuttan et al. 1989; Rao & Rao Secondary data collection was made from all the 1991; Apte 1998, 2014; Rao 2003; KSCSTE 2013), often published records on molluscan fauna of the islands and associated with a sandy bottom and -covered reefs. available checklists (Appukuttan et al. 1989; Rao & Rao Found in littoral and sublittoral zones, in tidal pools 1991; Apte 1998; Rao 2003; Ramakrishna & Dey 2010; and low levels to a depth of around 20m. This is Venkataraman et al. 2012; KSCSTE 2013; Apte 2014). A a common strombid in Lakshadweep. The flesh of this checklist of molluscs in various schedules of WPA of India species was earlier collected by the local people for the was prepared with updated names and classification. preparation of pickles. Included in Schedule IV of WPA of India. Not listed in CITES and not evaluated in IUCN Red List of Threatened Species. Currently the developmental RESULTS AND DISCUSSION stages are collected by the shell collectors without knowing its identity. Further, the juvenile stages of this The present survey recorded the presence of 14 species are almost impossible to distinguish from the species of molluscs representing the various schedules juveniles of Harpago arthriticus. of WPA of India. Of these, Placuna placenta (Linnaeus, 1758) (recorded from Kavaratti) and Tudicla spirillus 2. Harpago arthriticus (Röding, 1798) (Images 2 a-c) (Linnaeus, 1767) (recorded from Kalpeni) are new Apte 1998, p.32; Apte 2014, p. 59. records from Lakshadweep. A checklist of marine Description: Shell thick, robust, ovate and heavy molluscs in the various schedules of WPA and species with five labial digitations. turned to the recorded from Lakshadweep (Table 1) shows that 19 left. It looks similar to H. chiragra, but differs from the species of scheduled molluscs were recorded from the same with slightly depressed, broad rounded arching on island ecosystem. the end of aperture. This smaller species is with heavy whitish creamy plications in the aperture between which

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© Bijukumar & Ravinesh

a c e g

b d i j

k

Image 1. Linnaeus. f h a–h - various life stages; i - dorsal view of adult shell; j - ventral view of adult shell; k - lateral view of adult shell (scale = 1cm)

are dark purplish, streaks, which help distinguishing the from the juveniles of H. chiragra. species. Large “stromboid notch” located between the 4th and 5th labial digitations. The outer lip is yellowish 3. Lambis crocata (Link, 1807) (Images 3 a-c) or pinkish yellow with splotches of purplish underlying Apte 2014, p. 60. the white spiral lirae. Columella brownish-purple; four Description: Shape irregular with long, slender, spinal cords on the parietal wall. Peiostracum thin, curved siphonal digitations. The whorls are encircled by varnish like. Operculum chitinous, auburn brown colour, three rows of small knobs and many spiral threads. The fusiform, with fine striations on one side. spines on the outer lip, including the one enveloping the Remarks: In Lakshadweep it was recorded from posterior canal, are tubular in structure, but are usually Kalpeni (Apte 1998, 2014) and is rare in occurrence. filled up when the shell has fully grown, whereas only Included in Schedule IV of WPA. Not listed in CITES. Not the most anterior spine remains tubular as the anterior evaluated by IUCN Red List of Threatened Species. The canal. Anterior canal expanded in a very long and juvenile stages of this species are difficult to distinguish slender digitation. The outer surface is white in colour

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Table 1. List of marine molluscs in various Schedules of Wildlife (Protection) Act of India

Category in Records from Classification Common Name References WPA Lakshadweep Phylum: Mollusca Class: Gastropoda Subclass: Vetigastropoda 1. Top Shell Schedule IV Not recorded Superfamily: Trochoidae Family: Tegulidae Tectus niloticus (Linnaeus, 1767) Family: Turbinidae Great Green 2. Schedule IV Not recorded Turbo marmoratus Linnaeus, 1758 Turbon Subclass : Order: Recorded from Schedule IV 3. Family: Sibbald’s Lakshadweep; specific Apte 2014 plicata sibbaldi (G.B. locality not mentioned. Sowerby II, 1842) Found around all Appukuttan et al. (1989), Rao & Rao (1991), Chiragra Spider 4. Harpago chiragra (Linnaeus, 1758) Schedule IV inhabited islands of Apte (1998), Rao (2003), KSCSTE (2013), Apte Conch Lakshadweep (2014), Present study Arthritic Spider Rao & Rao (1991), Apte (1998), Rao (2003), 5. Harpago arthriticus (Röding, 1798) Schedule IV Kalpeni Conch KSCSTE (2013), Apte (2014), Present study Orange Spider Minicoy, Kalpeni, 6. Lambis crocata (Link, 1807) Schedule IV Apte 2014, Present study Conch Kavaratti Found around all Appukuttan et al. (1989), Rao & Rao (1991), Truncate spider 7. ([Lightfoot], 1786) Schedule IV inhabited islands of Apte (1998), Rao (2003), KSCSTE (2013), Apte Conch Lakshadweep 2014, Present study Milleped Spider 8. Lambis millepeda (Linnaeus, 1758) Schedule IV Not recorded Conch Recorded from Lambis indomaris Abbott, 1961[Listed 9. Scorpio Conch Schedule IV Lakshadweep; specific Apte (1998), Apte (2014), Present study as (Linnaeus, 1758)] locality not mentioned. Family: Minicoy, Kalpeni, Rao & Rao (1991), Apte (1998), Rao (2003), Horned Helmet Schedule I 10. cornuta (Linnaeus, 1758) Androth, Bangaram KSCSTE (2013), Apte (2014), Present study Minicoy, Kalpeni, Appukuttan et al. (1989), Rao & Rao (1991), Bull Mouth 11. rufa (Linnaeus, 1758) Schedule I Androth, Bangaram, Apte (1998), Rao (2003), KSCSTE (2013), Apte Helmet Chetlet (2014), Present study Appukuttan et al. (1989), Rao & Rao (1991), Family: Ranellidae Agatti, Amini, Chetlet, Trumpet Triton Schedule I Apte (1998), Rao (2003), KSCSTE (2013), Apte 12. tritonis (Linnaeus, 1758) Kalpeni 2014, Present study Recorded from Family: Limacina Schedule IV Lakshadweep; specific Apte (2014) 13 limacina (Lamarck, 1810) locality not mentioned. Recorded from mappa (Linnaeus, Schedule IV 14 Map cowrie Lakshadweep; specific Apte (2014) 1758) locality not mentioned. Appukuttan et al. (1989), Rao & Rao (1991), 15 talpa (Linnaeus, 1758) Mole Cowrie Schedule IV Kavaratti, Kalpeni Apte (1998), Rao (2003), KSCSTE (2013), Apte (2014), Present study Order: 16. Family: Turbinellidae Dog Chunk Schedule I Kalpeni, Bitra Present study Tudicla spirillus (Linnaeus, 1767) Family: 17. milneedwardsi Glory of India Schedule I Not recorded Jousseaume, 1894 Family: Fasciolariidae Recorded from Trapezium 18. Pleuroploca trapezium (Linnaeus, Schedule IV Lakshadweep; specific Apte (2014) Horse Conch 1758) locality not mentioned. Recorded from Family: 19. Schedule IV Lakshadweep; specific Apte (2014) arausiaca (Ligthfoot, 1786) Vaxillate Volute locality not mentioned. Class: Bivalvia Subclass: Pteriomorphia Order: Pectinoida Window Pane 20. Schedule IV Kavaratti Present study Family: Placunidae Oyster Placuna placenta (Linnaeus, 1758) Subclass: Heterodonta Order: Veneroida Found around all Appukuttan et al. (1989), Rao & Rao (1991), Elongate Giant 21. Superfamily: Cardioidea Schedule I inhabited islands of Apte (1998), Rao (2003), KSCSTE (2013), Clam Family: Cardiidae Lakshadweep Present study Tridacna maxima (Röding, 1798)

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Category in Records from Classification Common Name References WPA Lakshadweep Fluted Giant Agatti, Bangaram, Bitra, 22. Tridacna squamosa Lamarck, 1819 Schedule I Appukuttan et al. (1989), Present study Clam Chetlet 23. Hippopus hippopus (Linnaeus, 1758) Bear Paw Clam Schedule I Not Recorded Class: Cephalopoda Subclass: Nautiloidea Ramadoss (2003), Present study 24. Order: Nautilida Chambered Schedule I Bitra Family: Nautilidae Nautilus Nautilus pompilius Linnaeus, 1758

a b a b

c c & Ravinesh © Bijukumar © Bijukumar & Ravinesh © Bijukumar Image 2. Harpago arthriticus Röding Image 3. Lambis crocata Link a - dorsal view; b - ventral view; c - lateral view (scale = 1cm) a - dorsal view; b - ventral view; c - lateral view (scale = 1cm) and spotted with brown, but is covered over by a dark is extensively glazed. Columella and the outer lip of the brown . The interior of aperture is smooth aperture is nearly smooth. Outer lip with a well-marked and orange-red in colour. Operculum slightly curved, stromboid notch and six slender, hollow digitations. brown and its edges smooth. Inner lip with a heavy, extensive callus covering most of Shell length: 105–160 mm. the ventral side of the body and spire, including Remarks: This species was recorded from sometimes the in mature specimens. Siphonal Lakshadweep by Apte (2014). In the present study canal forming a wide and moderately developed specimens were recorded from Minicoy, Kalpeni, and digitation anteriorly slightly bent towards the right Kavaratti islands. Rare in occurrence. Specimens were (Image 4f). Operculum brown, chitinous, elongate and observed at shallow tidal pools and lagoons, around slightly curved. Outside of shell cream, sometimes 5–10 m depth. Included in Schedule IV of WPA. Not with spare light brown to tan speckles. Aperture white listed in CITES. Not evaluated in IUCN Red List of deep inside, outer lip and ventral callus creamy pink, Threatened Species. sometimes becoming purplish towards the edges of the glazed area. 4. Lambis truncata ([Lightfoot], 1786)(Images 4 a-i) In L. truncata the juvenile shells are long, with a thin Appukuttan et al. 1989, p.79; Venkataraman et al. outer lip, which is free from claws (Images 4 a,b). In later 2012, p. 191; KSCSTE 2013, p. 146; Apte 2014, p. 59. stages the outer lip flares up (Images 4 c,d) and then Description: Shells large and massive, ovate, with develops digitations (Images 4 e,f). In adults the outer spire very obtuse and knotty. Four uppermost whorls lips become thicker and slightly curve outwards in later are depressed, so as to give the spire a truncated stages (Images 4 h,i). Juveniles are easily distinguishable appearance, and each of the other whorls have a row as the apex is flat, hence the nametruncata. of strong rounded knobs. The ventral side of the shell Shell length: 320–380 mm.

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a c e

g h

b d f i

Image 4. Lambis truncata Lightfoot a–f - various life stages; g - dorsal view of adult shell; h - ventral view of adult shell; i - lateral view of adult shell (scale = 1cm)

Remarks: This species was recorded only from the shallow waters around all the inhabited islands of Lakshadweep (Appukuttan et al. 1989; KSCSTE 2013; Apte 2014). Common in appearance. They were found in areas with sandy bottom. Included in Schedule IV of WPA. Not listed in CITES. Not evaluated in IUCN Red List of Threatened Species.

5. Lambis indomaris Abbott, 1961(Images 5 a-c) Murnau, Germany © Ulrich Wieneke, Rao 2003, p. 149; Apte, 2014, p. 60 Description: Shell ovate, tuberculated, and transversely striated, with six knobbed labial digitations and a long slender knobbed siphonal digitation, which is a b strongly curved to the right. The lobe on the left side of the first digitation is either very much reduced or bent back around the apex of the shell. Digitations 4, 5 and 6 are more stunted. Whorls 9–11. Siphonal canal long, strongly curved to the right and bearing 4-5 weak knobs. Aperture rather narrow, quadrate and deep within its solid purple. c Shell greyish-white colour, slightly mottled with brown; throat saffron-coloured, and both the lips are Image 5. Lambis indomaris Abott a - ventral view; b - dorsal view; c - lateral view (scale = 1cm) purplish. Inside of outer lip with numerous crowded, white, slightly raised spiral lirae which end before they reach the broad, smooth yellow orange edge of the Remarks: Rao (2003) and Apte (2014) recorded this outer lip. Columella and parietal wall brownish-purple species from Lakshadweep, though the exact locality and overlaid with about 30–40 white to tan white, raised was not mentioned. We have not found this species spiral lirae. Operculum long, slightly curved, light brown, from any of the islands and it may be assumed that it is with 10 small serrations. very rare in occurrence in these islands. The population Shell length: 90–135 mm. status of the species needs to be studied in Lakshadweep.

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Included in Schedule IV of WPA and not listed in CITES. Included in Schedule I of WPA of India. Not listed in Not evaluated by IUCN Red List of Threatened Species, CITES. Not evaluated in IUCN red list. This species plays either. an important ecosystem function in the islands as the predator of crown-of-thorns starfish (Acanthaster 6. (Linnaeus, 1758) (Images 6 a-c) planci), which feeds on corals (Poutiers 1998). Rao & Rao 1991, p. 277; Rao 2003, p. 201; Venkataraman et al. 2012, p. 197; KSCSTE 2013, p. 144; 7. (Linnaeus, 1758) (Images 7 a-d) Apte, 2014, p. 87. Rao & Rao 1991, p. 277; Rao 2003, p. 202; Description: Shell very solid, heavy, rotund, with Venkataraman et al. 2012, p. 197; KSCSTE 2013, p large horn-like knobs and a wide flat base. Last whorl 146;Apte 2014, p. 88. with 3 or 4 spiral rows of large tubercles, those at Description: Shell thick and heavy, ovate with shoulder being much longer and stouter than the others. a strongly callous apertural side. Spire short, with Outer surface of whorls finely pitted, from numerous channelled sutures and fine nodulose spiral ribs, but intersecting axial and spiral threads. Aperture long and lacking axial varices. with rounded shoulder, narrow, heavily calloused. Outer lip thickened in a broad 3 or 4 spiral rows of rounded knobs posteriorly, two and flat shelf, with a dorsally recurved outer edge and spiral rows of axial ridges anteriorly, and with 2 or 3 with 5–7 strong teeth on its inner edge. Inner lip with an nodulose axial riblets between them. Knobs at shoulder extensive callous shield, forming a flange along left side not longer and stouter than the others. Aperture long of body whorl, and produced over the spire to join the and narrow, with an oblique furrow at posterior end. outer lip at its posterior end. Columella with irregular Outer lip very thick, produced over the spire, with spiral ridges. Operculum elongate-ovate, about one- strong teeth along the inner edge. Inner lip strongly fourth the length of aperture. lirate, with an extensive, thick and convex callous shield. Dorsal side and spire greyish-white, often somewhat Columellar margin with an axial swelling bearing strong spotted with light brown. Calloused ventral side glossy teeth. Operculum circular to oval, small, with concentric cream or orange, with two spiral rows of brown spots growth lines, about one-tenth the length of aperture. in the central region. Teeth and ridges of the aperture Dorsal side and spire orange-brown or reddish, white. Outer lip with 6 or 7 broad patches of brown on mottled and blotched with dark and light brown and its dorsal side and outer edge. grey. Callous ventral side glossy, creamy orange, Shell length: 50–410 mm. becoming deep red around and inside the aperture, and Remarks: Also known as “King Shell”, this largest stained with dark brown between the ridges of the inner of the helmet shells was recorded from the waters lip. Teeth and ridges of the aperture whitish. around Minicoy, Kalpeni, Androth and Bangaram Remarks: This species was recorded from the shallow islands of Lakshadweep (Rao & Rao 1991; Rao 2003; coastal waters of Minicoy, Kalpeni, Androth, Bangaram, Venkataraman et al. 2012; KSCSTE 2013; Apte 2014). and Chetlet islands (Rao & Rao 1991; Rao, 2003;

a b a b

c c d © Bijukumar & Ravinesh © Bijukumar © Bijukumar & Ravinesh © Bijukumar

Image 6. Cassis cornuta Linnaeus Image 7. Cypraecassis rufa Linnaeus a - spire; b - dorsal view; c - ventral view (scale = 1cm) a - lateral view; b - ventral view; c - dorsal view; d - spire (scale = 1cm)

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Venkataraman et al. 2012; KSCSTE 2013; Apte 2014). from some countries because of the absence of trade Included in Schedule I of WPA. Not listed in CITES. Not data. Not listed in IUCN Red List of Threatened Species . evaluated in IUCN Red List of Threatened Species. As the predator of crown-of-thorns Starfish (Acanthaster planci) which feeds on corals, this species plays a major 8. Charonia tritonis (Linnaeus, 1758) (Images 8 a, b) ecosystem function in the islands (Poutiers 1998). Rao & Rao 1991, p. 278; Ramakrishna & Dey 2003, p. 19; Rao 2003, p. 206; KSCSTE 2013,p. 144; Apte 2014, 9. (Linnaeus, 1758) (Images 9 a, b) p. 91. Rao & Rao 1991, p. 277; Apte 1998, p. 23; Rao 2003, Description: Spire very tall, occupying more than p. 174; KSCSTE 2013,p. 149; Apte 2014, p. 76. one-third of the total length. Broad bodywhorl with Description: Shell large, solid, almost cylindrical in two prominent varices. Deeply grooved flattened spinal outline, with widely convex to nearly straight lateral spiral ribs and occasional fine extra ribs below. Aperture margins and slightly produced anterior and posterior broader, bordered by the reflexed margin of inner lip extremities. Dorsal side moderately swollen, without a bearing many plaits and by the outer lip having a wavy groove. Ventral side flat, rounded laterally, with margin. Shell creamy pink with reddish or dark brown a narrow, straight aperture which is only a little curved crescents and patches. Aperture orange brown with posteriorly. Apertural teeth numerous, short and fine. white channels between teeth on outer lip. Teeth of inner lip not extending far into the aperture. Shell length: 390-450 mm. Anterior longitudinal furrow of inner lip short and rather Remarks: Recorded earlier from Agatti, Amini, deep. Chetlet, and Kalpeni islands (Rao & Rao 1991; Rao Colour pattern variable. The dorsal surface smooth 2003; Ramakrishna & Dey 2003, KSCSTE 2013; Apte and shiny, brown or yellowish-brown, with three or four 2014), this species is found among shallow coral seas yellow or light brown transverse bands. The margins, and on sandy bottoms. As a well-known predator base and teeth completely dark brown or black; of the crown-of-thorns starfish, it has a critical role in interstices of teeth stained with cream or white. In coral reef ecosystem. During the present survey none the living mantle is greyish or black, with long of the specimens could be located. Over exploitation, cylindrical papillae. especially for shell trade could be the reason for the Shell length: 80–110 mm. decline of this species in Lakshadweep. Included in Remarks: This species was earlier recorded from Schedule I of WPA. Not listed in CITES despite requests Kavaratti and Kalpeni islands (Rao & Rao 1991; Apte 1998; Rao 2003; KSCSTE 2013; Apte 2014). Included in Schedule I of WPA. Not listed in CITES. Not listed in IUCN Red List of Threatened Species. During the study, one © Bijukumar & Ravinesh © Bijukumar © Bijukumar & Ravinesh © Bijukumar

a b a b

Image 8. Charonia tritonis Linnaeus Image 9. Talparia talpa Linnaeus a - dorsal view; b - ventral view (scale = 1 cm) a - dorsal view; b - ventral view (scale = 1cm)

7262 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2015 | 7(6): 7253–7268 Molluscan fauna of Lakshadweep Bijukumar et al. specimen was recorded from Kalpeni. Detailed surveys Threatened Species. are required for assessing the status of this species in Lakshadweep, as the collection of cowries is one of 11. Placuna placenta (Linnaeus, 1758) (Images 11 a-c) the major economic activities of the local population. Description: Asymmetrical with a characteristically Distinguishing this species from the other species is flat, thin, translucent shell. The valves form an orbicular rather difficult in the field. outline, which is compressed laterally; the right is almost flat while the left is weakly convex. Outer 10.Tudicla spirillus (Linnaeus, 1767) (Images 10 a-c) shell surface marked by projecting imbricate lamellae. Description: Shell moderately solid, pyriform and with Inner shell surface smooth and polished. Generally a long, sinuous, smooth siphonal canal. Nuclear whorls translucent and colourless, growing more opaque and mammillate, projecting, swollen, pinkish or yellowish. white with age. Spire flat. Spiral of numerous, raised threads Shell length: 120mm. which are squarish and broad on the base of the whorls. Remarks: The present study marks the record of Last whorl with a squarish periphery, bounded above by this species for the first time from Kavaratti Island of a sharp, wavy keel and below by a series of 6–8 round, Lakshadweep. One specimen was observed on sandy low nodules. Aperture ovate-round, glossy, pinkish to bottom. Further surveys are required to assess the purple within, and bounded on the outer side by a sharp population status of this species included in schedule IV lip. Inside of outer wall with numerous, fine, raised, of WPA. Not in CITES and IUCN Red List of Threatened spiral ridges. Parietal shield well-developed, smooth Species . and glossy white. Posterior canal region with a swollen, button-like, white callus. Base of columella with a single, 12. Tridacna maxima (Röding, 1798) (Images 12 a-c) strong, spiral plica. Siphonal canal open along its length, Rao & Rao 1991, p. 286; KSCSTE 2013,p. 150 long and slightly sinuous. Colour of shell shiny cream to Description: Valves heavy and thick. Valve margins pinkish grey with sparse flecks of light-brown. undulate with about five generally sharply triangular Shell length: 70–85 mm. extremities of rib interstices. Hinge line less than half of Remarks: The present study records this species shell length. Inequilateral valves are elongate to short- for the first time from Kalpeni and Bitra islands of obtuse-triangular. Raised external valve sculpture. Large Lakshadweep. The recorded two specimens were byssal opening. Greyish-white shell tinted with yellow observed on sandy bottom. Further surveys are required or pinkish-orange. Bright blue, green or brown coloured to assess the population status of this species included mantle conceals the distinctively furrowed edges of in schedule IV of WPA. Not in CITES and IUCN Red List of shell.

a b

a b

c

c & Ravinesh © Bijukumar © Bijukumar & Ravinesh © Bijukumar Image 11. Placuna placenta Linnaeus Image 10. Tudicla spirillus (Linnaeus) a - dorsal shell; b- ventral shell; c- dorsal view of both shells opened a - dorsal view; b - ventral view; c - spire (scale = 1cm) (scale = 1cm)

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a a

b

b

c c

Image 13. Tridacna squamosa Lamarck a - exterior of right valve; b - interior of left valve; c - ventral view Image 12. Tridacna maxima Röding showing byssal orifice (scale = 1cm) a - exterior of right valve; b - interior of left valve; c - ventral view showing byssal orifice (scale = 1cm)

and conservation of T. maxima in the Lakshadweep Shell length: 250–350mm. Archipelago. Also known by the name ‘Small Giant Remarks: It is one of the widespread scheduled Clam’, this species grows on rocks and crevices of coral species recorded from almost all the islands of reefs, usually attached by the byssus threads. Included Lakshadweep (Rao & Rao 1991; KSCSTE 2013). Apte in schedule I of WPA. Not listed in CITES. Listed as (2013) reported on the ecology, population dynamics ‘Lower Risk’ in the IUCN Red List of Threatened Species

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(IUCN, 2013). Shell length: 200mm Remarks: This species is distributed in the Indo-West 13. Tridacna squamosa Lamarck, 1819 (Images 13 a-c) Pacific; the Philippines, via Indonesia and New Guinea Appukuttan et al. 1989, p. 83. to Fiji; north-eastern and north-western . In Description: Valves moderately thick and heavy. Shell India this species is reported only from Andaman and equilateral. Dorsal margin fan-shaped in outline. Valve Nicobar islands. During the present study one dead margins undulate with 4–6 pointed to bluntly rounded, specimen was recorded from Bitra Island, from a coral crenulated extremities of rib interstices. Hinge line half rock bottom. More research is warranted to establish of shell length. Characteristic broad leaf-like projecting whether this species actually lives around these islands. scales on primary folds of ribs. Small to medium sized byssal opening. Colour of shell tan or grey but new growth is often CONSERVATION white. Mantle colour patterns quite variable, brown, green, blue or purple coloured with large irregular The threats to biodiversity of Lakshadweep, as blotches. summarized in the National Biodiversity Strategy and Shell length: 300–400 mm. Action Plan (NBSAP), can be due to: (i) the fragility Remarks: Commonly referred to as scaly clam, of the island ecosystem, (ii) the imposing of locally fluted clam or giant clam, this species was recorded unsuitable development programmes which jeopardize from the Agatti, Bangaram, Bitra, and Chetlet islands by the economy and ecology of the islands, and (iii) the Appukuttan et al. (1989). We recorded this species from lack of knowledge about what was there creates a Kavaratti Island. Detailed field surveys are required to lacuna in understanding the depth of biodiversity loss relocate the populations around the various islands of (MoEF & Kalpavriksh 2004). As a coral island ecosystem, Lakshadweep. Included in Schedule I of WPA. Not listed the entire biological diversity is primarily dependent in CITES. Listed as ‘Lower Risk’ in the IUCN Red List of on corals. The pressure on the ecosystem as well as Threatened Species (IUCN 2013). biodiversity at present is mainly of anthropogenic origin. Global environmental issues such as climate change and 14. Nautilus pompilius Linnaeus, 1758 (Images 14 a, b) the pressure created by increasing human population Ramadoss 2003, p.207. and demand for natural resources create a host of Description: Large and thick shell, with smooth threats to the sensitive atoll islands, lagoon and the surface, polished and marked by reddish-brown wavy surrounding open ocean (KSCSTE 2013). Elevated Sea lines against creamy background. Parietal wall of Surface Temperatures (SSTs) caused by El Nino Southern aperture black. Shell spirally coiled and partitioned Oscillation (ENSO) and climate change have already internally into a series of gas-filled flotation chambers. reported to affect the health of the coral reef ecosystem The lives in the large external shell. Below the in some islands (Arthur 2000). hood pro trudes about 90 small suck erless ten ta cles. Be- Despite the fact that marine molluscs represent an neath, there is a fun nel con tain ing two sep a rate lobes. estimated 23% of all extant marine taxa, conservation of marine molluscs remains neglected globally and often information regarding the status of commercially © Bijukumar & Ravinesh exploited marine molluscs has been overshadowed by the collapse of finfish fishery (McManus 1997). Research into the conservation status of molluscs has so far failed to reflect this importance, with minimal inclusion on the Red List of World Conservation Union (IUCN) (Peters et al. 2013). Inclusion of 24 species of marine molluscs under various schedules of the WPA notwithstanding, the research and conservation on these species received little attention in policy documents. A majority of the molluscs listed in various schedules of WPA of a b India are preferred items in shell trade markets. High

Image 14. Nautilus pompilius Linnaeus market demand, coupled with a lack of awareness and a - dorsal view of shell; b - lateral view of shell (scale 1 cm) inadequate enforcement were found to be the major

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2015 | 7(6): 7253–7268 7265 Molluscan fauna of Lakshadweep Bijukumar et al. driving forces behind the illegal marine curio trade (John communities should also be ensured for planning and et al. 2012). decision making with regard to biodiversity. A lack of quantitative data on abundance of species, Currently, there are 31 MPAs in India represented ranges, habitat requirements, dispersal, and connectivity under 33 national parks and sanctuaries, covering an among populations (Chapman 1999) remains the major area of 627,121ha. The only MPA in Lakshadweep is handicap in assessing the conservation status of species Piiti Wildlife Sanctuary (1ha), included in the category II listed in various schedules of WPA. For example, the (Venkataraman et al. 2013). In Andaman and Nicobar extirpation of Giant Triton Charonia tritonis in the Indo- islands, however, there are 15 MPAs covering an area Pacific has been attributed to over harvesting (Moore & of 67,254ha (about 11% of the total area of MPAs Ndobe 2008). Although there is no organised shell trade in India). Lakshadweep has many untouched and at present in Lakshadweep, many of the species are undisturbed pristine reefs like Suheli par with 78.96km2 collected for food. Further, the developmental stages of of lagoon area, Baliyapani par with 57.46km2 of lagoon scheduled molluscs such Harpago chiragra, H. arthritica area, Cheriyapani par with 172.59km2 of lagoon area and Lambis spp. are often difficult to identify in the field, and Perumul par with 83.02km2 of lagoon area, which which indirectly facilitate indiscriminate collection of remains to be studied fully for realising the magnitude these species. Many species of Strombidae, to which of biodiversity. While in the Indian context MPAs are the genera Harpago and Lambis belong, are eaten, and not a specific legal category, national parks, sanctuaries sometimes an important source of protein. This also and community conservation reserves declared either in reiterates the need for creating more awareness among selected biodiverse islands or declaration of a biosphere the islanders regarding the scheduled species of molluscs. reserve covering the whole Lakshadweep Archipelago is A poster on sea shells of Lakshadweep protected under strongly recommended for conservation and sustainable various schedules of Wildlife (Protection) Act of India management of biodiversity for future generations. has been prepared for creating awareness (Appendix 1). According to Peters et al. (2013) research is needed to assess the threat from rare shell collecting towards REFERENCES mollusc population decline to determine what measures should be taken to enable this activity to continue Aditi, N. & D.A. Apte (2013). Artisanal octopus fishery: socio- economics and management, pp 409–418. In: Venkataraman, K., sustainably while at the same time allowing for the C. Sivaperuman & C. Raghunathan (eds). Ecology and Conservation protection of vulnerable species. While Indian WPA of Tropical Marine Faunal Communities. Springer Berlin Heidelberg, prevents collection of included in various 481pp. Appukuttan, K.K., A. Chellam, A., K. Ramdoss, A.C.C. Victor & M.M. schedules, research priority should also focus on other Meiyappan (1989). Molluscan resources, pp. 77–92. In: Suseelan, C. molluscs such as cowries and octopuses which are (ed.). Marine Living Resources of the Union Territory of Lakshadweep: currently collected in large numbers. According to An Indicative Survey with Suggestions for Development. Bulletin, 43, Central Marine Fisheries Research Institute, Kochi, Kerala, 256pp. Apte et al. (2010) ever increasing economic interactions Apte, D.A. (1998). The Book of Indian Shells. Oxford University Press, between the densely populated island communities and Mumbai, 115pp. Apte, D.A. (2009). Opisthobranch fauna of Lakshadweep Islands, India the adjoining coral reefs of Lakshadweep can indirectly with 52 new records to Lakshadweep and 40 new records to India. affect the scheduled Giant Clam (Tridaca spp.), rendering Part 1. Journal of the Bombay Natural History Society 106: 162–175. population monitoring imperative. Apte, D.A. (2012a). Field Guide to the Marine Life of India - 1st Edition. Stusa Mudra Private Limited Mumbai, 502pp. Islands are very sensitive ecosystems. Hence Apte, D.A. (2012b). Ecology, population dynamics and conservation of strategies should be adopted to integrate biodiversity the Giant Clam Tridacna maxima (Röding, 1798) in the Lakshadweep into every developmental plan, especially in the context Archipelago. PhD Thesis, University of Mumbai, 183pp. Apte, D.A. (2014). Sea Shells of India. An Illustrated Guide to Common of rapid economic development of Lakshadweep. Gastropods. Bombay Natural History Society & Oxford University Preparation of people’s biodiversity register for the island Press, Mumbai, 197pp. ecosystems, as envisaged in the Biological Diversity Act Apte, D.A. & D. Sutirtha (2010).Ecological determinants and stochastic fluctuations of Tridacna maxima survival rate in Lakshadweep of India, would help in developing suitable management Archipelago. Systematic and Biodiversity 8(4): 461–469; http:// plans at the grass roots level. As each island is a separate dx.doi.org/10.1080/14772000.2010.537393 entity in terms of its biodiversity and pressures onit, Apte, D.A., D. Sutirtha & K.K. 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Coral Reefs of India - Status, Threats and Conservation Measures. Poutiers, J.M. (1998). Gastropods, pp. 363–648. In: Carpenter, K.E. IUCN, 305pp. & V.H. Niem (ed.). FAO Species Identification Guide for Fishery Apte, D.A. & B. Vishal (2014). New records of opisthobranchs from Purposes. The Living Marine Resources of the Western Central Lakshadweep, India (Mollusca: Heterobranchia). Journal of Pacific. Volume 1. Seaweeds, corals, bivalves, and gastropods. Threatened Taxa 6(3): 5562–5568; http://dx.doi.org/10.11609/ Rome, FAO. JoTT.o3487.5562-8. Prabhakaran, M.P., P.R. Jayachandran & S. Bijoynandan (2012). New Apte, D.A., A. Kulkarni & D. Sutirtha (2014). Giant Clam Tridacna record of Scapharca cornea (Bivalvia: Pteriomorpha: Arcidae) from maxima from Lakshadweep Archipelago, India: Understanding Minicoy Lagoon, Lakshadweep, India. Current Science 102 (11): Population Ecology. LAP Lambert Academic Publishing, Germany, 1516- 1518. 308pp. Prashanth, M. & K. Veenakumari (1996). 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Current status of Indian opisthobranch Ramachandran, A. & A. Joseph (eds.). Marine Biodiversity: Status, fauna, pp. 63–79. In: Venkataraman, K., C. Sivaperuman & C. Opportunities and Challenges. Department of Marine Biology, Raghunathan (eds). Ecology and Conservation of Tropical Marine Microbiology and Biochemistry, Cochin University of Science and Faunal Communities. Springer-Verlag Berlin Heidelberg, 481pp. Technology, Kerala. Moore, A. & S. Ndobe (2008). Reefs at risk in Central Sulawesi, Indonesia - status and Outlook. Proceedings of the 11th International Coral Reef Symposium, Ft. Lauderdale, Florida, 7–11 July 2008, pp. Author Details: Dr. A. Bijukumar is the Associate Professor and Head of the 840–844. Department of Aquatic Biology and Fisheries, University of Kerala and is Namboodiri, P.N. & P. Sivadas (1979). Zonation of molluscan specialized in taxonomy and biodiversity documentation. Mr. R. Ravinesh and assemblage at Kavaratti Atoll (Lacccadives). Mahasagar 12 (2): 239- Ms. A.R. Arathi are research scholars working in Department of Aquatic Biology 246. and Fisheries, University of Kerala and pursuing their doctoral work on molluscan Olson, D.M. & E. Dinerstein (1998). The global 200: A representation fauna of Lakshadweep and bivalves of west coast of India and Lakshadweep approach to conserving Earth’s most biologically valuable respectively. Dr. K.K. Idreesbabu is the Scientist, working with Department of ecoregions. Conservation Biology 12: 502–515. Science and Technology, Kavaratti, Union Territory of Lakshadweep. Peters, H., B.C. O’Leary, J.P. Hawkins, K.E. Carpenter & C.M. Roberts Author Contribution: ABK was the Principal Investigator of the project and was (2013). Conus: First comprehensive conservation red list assessment involved in taxonomic identification and preparation of the paper. RR, ARA and of a marine gastropod mollusc . PLoS ONE 8(12): e83353; KKI carried out field work in all the islands, identified the species in collaboration http://dx.doi.org/10.1371/journal.pone.0083353 with experts and assisted preparation of the manuscript.

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Appendix 1. Poster on sea shells of Lakshadweep protected under Wildlife (Protection) Act of India

Threatened Taxa 7268 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2015 | 7(6): 7253–7268